Phlogites Temporal range: | |
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Restoration of Phlogites (with tentacles extended) Phlogites longus fossil | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Superphylum: | Deuterostomia |
Stem group: | Ambulacraria |
Clade: | † Cambroernida |
Genus: | † Phlogites Luo & Hu, 1999 |
Species: | †P. longus |
Binomial name | |
†Phlogites longus Luo & Hu, 1999 | |
Synonyms | |
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Phlogites is a member of the extinct ambulacrarian stem group Cambroernida, occupying an intermediate position between the basal Herpetogaster and the more derived Eldonioidea. [4] [5] It is known from the Lower Cambrian Haikou Chengjiang deposits of China. [4]
Phlogites was a cup-shaped animal with a branching tentacular feeding system leading to a dextrally coiled gut with a lateral anus. Sources differ as to the number of tentacles, with two, [1] [5] three, [7] three or five, [2] or four or five [8] tentacles said to be on the anterior part of the calyx. Smooth semi-circular lobes are present between the tentacles. [8]
Compared to the earlier-diverging cambroernid Herpetogaster, Phlogites lacked segmentation and had a more massive stolon that is contiguous with the body. The coiling present in the external form of Herpetogaster became internal, except for a small lobate extension with the anus opening laterally. The tentacles of Phlogites were also more massive, and seem to have branched dichotomously. [8]
The following cladogram is simpllified from Li et al. 2023: [9]
Phlogites was initially considered to be an early lophophorate. [10] A new phylum, Dendrobrachia (Hou, Bergstrom, Ma and Zhao, 2006) was later proposed to encompass it. While some similarities were noted with the eldonioids (and phoronids), the authors were not sufficiently confident to either group Phlogites with the phoronids or include the eldonioids in the Dendrobrachia. [1]
Since ICZN precedence rules do not apply at the phylum level, later researchers opted to propose cambroernids as an informal group rather than expanding the phylum Dendrobrachia, noting that "not only is the concept of phyla essentialist but it serves to place problematic taxa in a phylogenetic limbo, rendering them effectively immune to further evolutionary analysis." [11] The informal group "cambroernids" was later given the formal name Cambroernida once further evidence was found to support its monophyly. [12]
The interpretation of Phlogites has been complicated by its proposed equation (as a senior synonym) [2] with the possible first-disovered Cambrian tunicate Cheungkongella ancestralis [13] (sometimes misspelt Cheungkungella) [14] by the discoverers of another candidate for the title of the oldest known Cambrian tunicate, Shankouclava . [15]
The discoverers of Shankouclava agreed with the discovers of Phlogites that Phlogites is most likely a lophophorate, although they noted possible affinities with ambulacrarians or with Sipuncula. [2] A different group of researchers agreed with synonymizing Cheungkongella with Phlogites, but proposed a new phylum, Dendrobrachia, with affinities to the Gnathifera. [1]
Neither group proposed tunicate affinities for Phlogites, but later works by the discoverer of Shankouclava contain statements such as "although Lower Cambrian Phlogites (=Cheungkongella) was claimed as a tunicate (Shu et al., 2001)" [16] that have led to confusion on this point [17] by associating the tunicate affinity claims that only ever applied to Cheungkongella with Phlogites. [18]
A researcher who was not part of any of the teams discovering Phlogites, Cheungkongella, or Shankouclava, or proposing Dendrobrachia, examined the fossils and concluded that the similarity between Phlogites and Cheungkongella was "superficial." [3] The question of whether Cheungkongella is a synonym of Phlogites remains unresolved, [15] but does not impact the current theory of Phlogites as a cambroernid. [19]
Vetulicolia is a group of bilaterian marine animals encompassing several extinct species from the Cambrian, and possibly Ediacaran, periods. As of 2023, the majority of workers favor placing Vetulicolians in the stem group of the Chordata, but some continue to favor a more crownward placement as a sister group to the Tunicata. It was initially erected as a monophyletic clade with the rank of phylum in 2001, with subsequent work supporting its monophyly. However, more recent research suggests that vetulicolians may be paraphyletic and form a basal evolutionary grade of stem chordates.
The Maotianshan Shales (帽天山页岩) are a series of Early Cambrian sedimentary deposits in the Chiungchussu Formation, famous for their Konservat Lagerstätten, deposits known for the exceptional preservation of fossilized organisms or traces. The Maotianshan Shales form one of some forty Cambrian fossil locations worldwide exhibiting exquisite preservation of rarely preserved, non-mineralized soft tissue, comparable to the fossils of the Burgess Shale of British Columbia, Canada. They take their name from Maotianshan Hill in Chengjiang County, Yunnan Province, China.
Yunnanozoon lividum is an extinct species of bilaterian animal from the Lower Cambrian Chengjiang biota of Yunnan province, China. Its affinities have been long the subject of controversy.
Vetulicola is an extinct genus of marine animal discovered from the Cambrian of China. It is the eponymous member of the enigmatic phylum Vetulicolia, which is of uncertain affinities but may belong to the deuterostomes. The name was derived from Vetulicola cuneata, the first species described by Hou Xian-guang in 1987 from the Lower Cambrian Chiungchussu Formation in Chengjiang, China.
Didazoon haoae is an extinct species of vetulicolid vetulicolian described by Shu, et al. based on fossils found in the Qiongzhusi (Chiungchussu) Formation, Yu'anshan Member, Lower Cambrian, in the Dabanqiao area (Kunming), about 60 km northwest of Chengjiang, China.
Pomatrum is an extinct vetulicolian, the senior synonym of Xidazoon; the latter taxon was described by Shu, et al. (1999) based on fossils found in the Qiongzhusi (Chiungchussu) Formation, Yu'anshan Member, Lower Cambrian, Haikou, (Kunming), about 50 km west of Chengjiang, China.
Vetulicolidae is a vetulicolian family from the Cambrian Stage 3 Maotianshan Shale and Sirius Passet Lagerstätte that consists of Vetulicola, Beidazoon, and Ooedigera. It is distinguished from the Didazoonidae by a harder body wall and the lack of an oral disc.
Didazoonidae is a vetulicolian family within the order Vetulicolata. It is characterized by a relatively thin-walled, non-biomineralized body and a large, round anterior opening surrounded by an oral disc. It may be paraphyletic, even if the phylum Vetulicolia is monophyletic.
Deuterostomes are bilaterian animals of the superphylum Deuterostomia, typically characterized by their anus forming before the mouth during embryonic development. Deuterostomia is further divided into four phyla: Chordata, Echinodermata, Hemichordata, and the extinct Vetulicolia known from Cambrian fossils. The extinct clade Cambroernida is thought to be a member of Deuterostomia.
Archaeopriapulida is a group of priapulid worms known from Cambrian lagerstätte. The group is closely related to, and very similar to, the modern Priapulids. It is unclear whether it is mono- or polyphyletic. Despite a remarkable morphological similarity to their modern cousins, they fall outside of the priapulid crown group, which is not unambiguously represented in the fossil record until the Carboniferous. In addition to well-preserved body fossils, remains of several archaeopriapulid taxa are known to have been preserved primarily as organic microfossils, such as isolated scalids and pharyngeal teeth. They are probably closely related or paraphyletic to the palaeoscolecids; the relationship between these basal worms is somewhat unresolved.
Eldonia is an extinct soft-bodied cambroernid best known from the Fossil Ridge outcrops of the Burgess Shale, particularly in the 'Great Eldonia layer' in the Walcott Quarry. In addition to over 550 specimens collected by Walcott, 224 specimens of Eldonia are known from the Greater Phyllopod bed, where they comprise 0.43% of the community. Species also occur in the Chengjiang biota, Siberia, and in Upper Ordovician strata of Morocco.
Herpetogaster is an extinct cambroernid genus of animal from the Early Cambrian Chengjiang biota of China, Blang Formation of China, Pioche Formation of Nevada and Middle Cambrian Burgess Shale of Canada containing the species Herpetogaster collinsi and Herpetogaster haiyanensis.
Eldonioids are an extinct clade of disc-shaped cambroernids, the Eldonioidea, which lived in the early to middle Paleozoic. The terms "eldonioid" and "eldoniid" have been used somewhat informally and interchangeably, but technically refer to members of the class Eldonioidea and the family Eldoniidae, respectively. The lifestyle of eldonioids is still an unresolved question; some authors reconstruct eldonioids as free-floating planktonic predators similar to jellyfish, while others argue that they were passive detritivores, embedded within the seabed for much of their life.
Rotadiscus is a genus of discoidal animal known from the Cambrian Chengjiang biota and classified with the eldonioids.
The Cambroernida are a clade of unusual Paleozoic animals with coiled bodies and filamentous tentacles. They include a number of early to middle Paleozoic genera noted as "bizarre" or "orphan" taxa, meaning that their affinities with other animals, living or extinct, have long been uncertain. While initially defined as an "informal stem group," later work with better-preserved fossils has strengthened the argument for Cambroernida as a monophyletic clade.
Cotyledion tylodes is an extinct, stalked filter-feeder known from the Chengjiang lagerstatten. The living animal reached a couple of centimetres in height, and bore a loose scleritome of ovoid sclerites. Its interpretation has been controversial, but it is currently thought to be a member of the Entoprocta stem group.
Vetulicola rectangulata is a species of extinct animal from the Early Cambrian of the Chengjiang biota of China. Regarded as a deuterostome, it has characteristic rectangular anterior body on which the posterior tail region is attached. It was described by Luo Huilin and Hu Shi-xue in 1999.
Shankouclava is an extinct genus of tunicates. It is one of two candidates for the oldest member of this group, dating to 518 million years ago. It has been found in the Lower Cambrian Maotianshan Shale at Shankou village, Anning, near Kunming. Each of the eight specimens found and used for description were isolated, suggesting that the genus was solitary and not colonial.
Banfiidae is a family of extinct banffozoan animals from North America and China. The family name is sometimes spelt Banffidae. It includes Banffia, Heteromorphus, and possibly Skeemella. The family may be paraphyletic. The family may be paraphyletic. A Heteromorphus-like dwarf "Form A" is allied with this group at the class level, but has not been formally described or assigned to this family.
Cheungkongella is a fossil organism from the lower Cambrian Chengjiang lagerstatte, the affinity of which has been the subject of debate. It was announced as a "probable" tunicate while noting the lack of definitive Cambrian fossils from that group. However, this affinity was later disputed in a paper announcing the discovery of Shankouclava, also from Chengjiang, as the oldest known tunicate. Cheungkongella has been accepted as a distinct taxon and possible tunicate by multiple workers not involved in its discovery, but the dispute remains unresolved.